Pollination

Pollination Definition: transfer of pollen from stamen to stigma Contrast with herbivory. Animals eating plant products but: 1) Most pollinators winged 2) If insects, adults involved directly and not larvae 3) Few mammals involved (except bats)

The flower Parts (see Fig. 6-1 in text)

Life Cycle Example of sporic meiosis: Gametophytes make gametes BY MITOSIS Zygote grows into diploid individual called sporophyte Sporophyte makes meiospores BY MEIOSIS Two bodies in one cycle: alternation of generations

Life Cycle Overview:

Life Cycle Overview: Fertilization: union of sperm with egg to form zygote

Floral variation Parts may be fused Example, petals fused to each other Like parts fused: connation (ex., petals to petals) Unlike parts fused: adnation (ex., stamens to petals). Snapdragon flower

Floral variation Fusing of petals can form floral tube (nectar made at bottom) Only long-tongued pollinators can reach it. Anisacanthus (Acanthaceae) flower

Floral variation Flowers with both stamens and pistils: perfect flowers Another kind of perfection….

Floral variation Flowers with both stamens and pistils: perfect flowers Some flowers imperfect. Either pistillate (have pistil) or staminate (have stamens). Pistillate flowers of Sagittaria Staminate flowers of Sagittaria

Floral variation Note: some species make pistillate flowers and carpellate flowers on separate individuals This termed dioecy (MUST outcross to reproduce sexually) Monoecy is when both sexes on same individual.

Floral variation Example of dioecious species: Persimmon (Diospyros) Persimmon fruits Example of dioecious species: Persimmon (Diospyros) Pistillate flower Staminate flower

Floral variation Some flowers are missing one or more sets of basic parts: incomplete flowers Note that all imperfect flowers are therefore incomplete!

Floral variation Floral symmetry: Radial: can be divided into similar halves by several planes Bilateral: can be divided into mirror images by 1 plane.

Floral variation Ovary position Superior ovary: other parts attach below ovary (hypogynous: “hypo-” =below, “gyn-” =female)

Floral variation Example of superior ovary in a lily flower (ovary is E)

Floral variation Ovary position Perigynous flower: ovary superior, but cup formed of fused sepals, petals, stamens around it.

Floral variation Ovary position Inferior ovary: other parts attach above ovary (epigynous: “epi-”=above, “gyn-”=female).

Floral variation Example of inferior ovary: squash flower (this one is pistillate). Ovary

Floral variation ray flowers disk flowers Some flowers assembled into groups of flowers: inflorescence Special inflorescence type: head Example, sunflower and its relatives Ray flowers have large fused petals (corollas fused), disk flowers small and crowded. ray flowers disk flowers

Floral variation Flowering dogwood (Cornus florida) Inflorescence: white structures are modified leaves (bracts) that act like petals. Closeup showing individual greenish flowers Inflorescence

Pollination Why flowers so varied? Many form mutualism with animals to achieve pollination

Mutualism Exceptions Some flowering plants are wind pollinated (anemophily) Some are water pollinated (hydrophily) Small, greenish grass flowers

Pollination as Mutualism Most flowering plants are pollinated by animals This usually viewed as mutualism (where both species benefit) Plant gets pollen transferred Animal gets “reward”

Rewards Pollen: high in protein Also has lipids, minerals, starch Can be renewed by: sequential anther dehiscence (multiple stamens) poricidal anthers (buzz pollination)

Rewards Nectar: sugary fluid produced by nectar glands (nectaries) in flower 10-60% mono- or disaccharides May have amino acids too (butterfly flowers) Renewable reward!

Rewards Oils/Resins: some used as construction materials, “cologne” (male solitary bee uses oil as female attractant), food for larvae (Krameria) Edible petals (pineapple guava: New Zealand) Krameria wax gland: wasp food!

Pollination Benefits of animal pollination for plant 1) Directed dispersal of pollen. Can get delivered from stamen to stigma with less waste Floral cues and attractants: Color and shape Scent Warmth (thermogenic plants: rare) Skunk cabbage

Pollination Benefits of animal pollination for plant 1) Directed dispersal of pollen. This aided by learning of floral visitors: decreases “handling time” Fosters “floral constancy” (visiting single species on foraging trip) by bees Bumblebee visits to touch-me-not

Pollination Benefits of animal pollination for plant stigma style 2) Style of flower as “selective racetrack” stigma style ovary 2 ovules

Pollination Benefits of animal pollination stigma style ovary 2 ovules 2) Style of flower as “selective racetrack” Keep in mind that 1 pollen grain can fertilize 1 ovule Suppose 5 pollen grains arrive on stigma Pollen grains stigma style ovary 2 ovules

Pollination Benefits of animal pollination stigma style ovary 2 ovules 2) Style of flower as “selective racetrack” Keep in mind that 1 pollen grain can fertilize 1 ovule Suppose 5 pollen grains arrive on stigma Start to make pollen tubes How many can fertilize an ovule? Pollen grains stigma pollen tubes style ovary 2 ovules

Pollination Benefits of animal pollination stigma style ovary 2 ovules 2) Style of flower as “selective racetrack” Keep in mind that 1 pollen grain can fertilize 1 ovule Suppose 5 pollen grains arrive on stigma Start to make pollen tubes How many can fertilize an ovule? 2! First 2 to arrive! Rest? LOSERS! Pollen grains stigma style ovary 2 ovules

Pollination Benefits of animal pollination stigma style ovary 2 ovules 2) Style of flower as “selective racetrack” Pollen tubes are haploid (1n) Haploid means only 1 allele (gene version) for every trait Pollen grains stigma style ovary 2 ovules

Pollination Benefits of animal pollination stigma style ovary 2 ovules 2) Style of flower as “selective racetrack” Pollen tubes are haploid (1n) Haploid means only 1 allele (gene version) for every trait If an allele is recessive, then it will be expressed (can’t be masked by another, dominant allele) Pollen grains stigma style ovary 2 ovules

Pollination Benefits of animal pollination stigma style ovary 2 ovules 2) Style of flower as “selective racetrack” Pollen tubes are haploid (1n) Haploid means only 1 allele (gene version) for every trait If an allele is recessive, then it will be expressed (can’t be masked by another, dominant allele) So, fittest (fastest) pollen grains mate Inferior genes don’t get passed to offspring. Pollen grains stigma style ovary 2 ovules

Pollination Style of flower as “selective racetrack” Is there evidence that this works? Example, Coyote melon Gourd growing in U.S. deserts.

Pollination Style of flower as “selective racetrack” Study done in 2000 showed that 1) takes 900 pollen grains to fully pollinate flower 2) 1 pollinator visit puts 650 grains/flower. By 2 hours, >4000 grains deposited on stigma 3) Seeds produced from over-pollinated flowers produced more vigorous seedlings (compared to seeds from flowers with <900 pollen grains on stigma).

Outcrossing Major benefit of sexual reproduction: generate genetic variation This enhanced by mating with others (outcrossing)

Outcrossing How to favor outcrossing: 1) Dioecy Bet hedging! Some selfing can occur if no cross pollination Imperfect flowers

Outcrossing How to favor outcrossing: 2) Floral morphology Heterostyly: Distyly and tristyly

Outcrossing How to favor outcrossing: 3) Floral phenology Protandry How to favor outcrossing: 3) Floral phenology Protandry: anthers dehisce first Protogyny: stigma becomes receptive first

Outcrossing How to favor outcrossing: 4) Self-incompatibility: prevents germination of self pollen or slows self pollen tube growth

Pollinator Specialization May be learned May be species-specific Monolecty: Flowers of 1 plant species visited Oligolecty: Flowers of few plant species visited Polylecty: Flowers of many plant species visited

Specialization Benefits of taxonomic specialization Better service: can match phenology of plant/pollinator Decrease competition (must match flower/pollinator traits) Plant: Minimize stigma clogging with heterospecific pollen

Pollination syndromes Disclaimer: Note that we must make broad generalizations and exceptions exist to most statements!

Pollination syndromes Bee pollination: Melittophily Bees: intelligent, agile visual animals: good eyesight (including UV light) good smellers (good sense of smell) day-active

Pollination syndromes Bee pollination Bee pollinated flowers: Colorful (usually not red) Landing platform: place where bee can land on flower Mildly fragrant. Nectar guides: patterns of lines or dots that can guide bee to reward Sometimes these only visible in UV light (which bees see). Petals in visible light (left) and UV (rt)

Pollination syndromes Fremontodendron decumbens (endangered chaparral shrub) flowers UV reflectance photo (right)

Pollination syndromes Beetle pollination: Cantharophily Beetles are: Clumsy Have poor vision Dumb(er) Active during the day (many flower-visiting ones).

Pollination syndromes Beetle pollination Beetle pollinated flowers are: Relatively large or grouped into large inflorescences Light colored Smelly (fruity or spicy smell) Inflorescence of Xanthosoma with beetle from it Dogwood inflorescence

Pollination syndromes Bird pollination: Ornithophily Birds are: agile long-beaked visual: see red colors well poor “smellers” Hummingbirds (native to Americas) can hover!

Pollination syndromes Bird-pollinated flowers are: red or orange have nectar hidden by long floral tube little or no fragrance no landing platform Anisacanthus (Acanthaceae) flower Aloe

Pollination syndromes Bird pollination: not always by hummingbirds Other birds from other areas: Sunbirds (Africa/Asia) Honeycreepers (Hawai’i). Golden-winged sunbird Iiwi (a honeycreeper) Crested honeycreeper

Pollination syndromes Butterfly pollination: Psychophily Butterflies have: good vision, good sense of smell, long coiled tongue. Must land on flower to visit it (can’t hover). coiled tongue tongue extended

Pollination syndromes Butterfly-pollinated flowers: Color varies (blue, yellow, orange) Landing platform present Nectar at bottom of floral tube Plumbago flowers Phlox flowers

Pollination syndromes Fly pollination: Myophily Flies have good sense of smell, especially flesh flies Attracted to rotting meat (lay eggs in meat, larvae are maggots).

Pollination syndromes Fly-pollinated flowers: Sapromyophily Smell like rotting meat Look like rotting meat (dark red, purple) Offer no reward: flies fooled by flower. Stapelia flower

Fly flower story: Rafflesia Ex, Rafflesia of Sumatra Root parasite Flower is only above-ground part.

Fly flower story: Rafflesia Ex, Rafflesia of Sumatra Largest single flower on Earth.

Pollination syndromes Fly pollination Largest inflorescence is made by fly-pollinated plant Corpse-flower. Inside view

Pollination syndromes Bat pollination: Chiropterophily Bats are flying mammals Nocturnal Eyesight good but echolocate Good sense of smell Agile, can hover when visiting flower.

Pollination syndromes Bat pollination Bat-pollinated flowers Open at night Lots of pollen and nectar as rewards White or light-colored Fragrant (sweet) May be pendant (hang from branches). Parkia flowers

Pollination syndromes Bat pollination Bat-pollinated flowers May also be made on tree trunks (tropics): Cauliflory Ex, cannonball tree of South America

Pollination syndromes Bat pollination: Mainly a tropical phenomenon In U.S., saguaro cactus is one of few bat-pollinated species. Saguaro cactus Saguaro flowers

Pollination syndromes Moth pollination: Phalaenophily Moths have: Poor vision (nocturnal) Excellent sense of smell Long coiled tongue.

Pollination syndromes Moth pollination Moths: Some (hawkmoths) can hover when visiting flowers. Hawkmoth

Pollination syndromes Moth pollination Moth-pollinated flowers: Open at night Sweet fragrance White or light-colored Nectar in tube.

Pollination syndromes Interesting moth story: nectar spur Nectar spur is long pouch, at bottom of which is nectar Moth uses long tongue to reach nectar Nectar spurs on columbine

Pollination syndromes Interesting moth story: nectar spur Nectar spur is long pouch, at bottom of which is nectar Moth uses long tongue to reach nectar Only if tube is longer than tongue will moth have to push into flower far enough to pick up pollen So, long spurred flowers reproduce better.

Pollination syndromes Which leads to longer moth tongues to reach all of the nectar in the longer tubes Which leads to longer tubes......

Pollination syndromes Which leads to longer moth tongues to reach all of the nectar in the longer tubes Which leads to longer tubes...... Some moth-pollinated orchids with long nectar spurs (almost one foot long!) Moth has extremely long tongue!

Pollination syndromes Moth pollination Angraecum orchid from Madagascar. Nectar spur almost 1 foot long!

Pollination syndromes Can read more in magazine for Gene Simmons fans. Gene Simmons from KISS

Sexual Mimics Flowers that mimic female bees or wasps Look like females Smell like females: chemical mimicry. One study showed flower more attractive than real female!! Sexual mimic orchids

Pollination syndromes Sexual mimics Males attempt to mate (pseudocopulation), pick up pollen, then fly to another flower and repeat process No reward supplied!. A male wasp “mating” with an Ophrys flower (how embarrassing...)

Pollination syndromes Another trick flower Grass pink orchid (Calopogon): grows in pitcher plant bogs in SE US Makes fake stamens on petal. Plant ecology class in bog Cattleya orchid (flower made right side up) Calopogon orchid (flower made upside down!)